1. M343 defines R1b in the broadest sense
2. P25 defines R1b1, making up most of R1b, and is often used to test for R1b.
It has been confused with m25, which is the marker for Q1a2 3. In some cases, major downstream mutations such as M269 are used to identify R1b, especially in regional or out-of-date studies

Haplogroup R1b (Y-DNA) is the dominant paternal lineage of Western Europe. In human genetics, Haplogroup R1b is the most frequently occurring Y-chromosomehaplogroup in Western Europe and in parts of sub-Saharan Central Africa (for example around Chad and Cameroon). R1b is present at lower frequencies throughout Eastern Europe, Western Asia, Central Asia, and parts of North Africa, South Asia, and Siberia. Due to European emigration it reaches high frequencies in the Americas and Australia. While Western Europe is dominated by the R1b1a2 (R-M269) branch of R1b, the mostly Chadic-speaking area in Africa is dominated by the branch known as R1b1c (R-V88). These represent two very successful "twigs" on a much bigger "family tree".

"R1b", "R1b1", and so on are "phylogenetic" or family tree based names which explain the branching of the family tree of R1b. For example R1b1a and R1b1b would be branches of R1b1, descending from a common ancestor. This means that these names can change with new discoveries.

The alternative way of naming haplogroups is to refer to the SNP mutations used to define and identify them, for example "R-M343" which is equivalent to "R1b." Haplogroup R1b is in other words now identified by the presence of the single-nucleotide polymorphism (SNP) mutation M343, which was discovered in 2004.[4] From 2002 to 2005, R1b was defined by the presence of the SNP named P25.

Standardized naming as described above, both using phylogenetic or mutational systems, was first proposed in 2002 by the Y Chromosome Consortium. Before 2002, today's Haplogroup R1b had a number of names in differing nomenclature systems, such as Hg1 and Eu18.[5]

After 2002, a major update of the YCC phylogenetic nomenclature was made in 2008 by Karafet et al. which took account of newer discoveries of branches which could be clearly defined by SNP mutations, including some which changed the understanding of R1b's family tree.[1] Since 2008 it has become increasing necessary to refer to the frequently updated listing made on the ISOGG website.[2]

Before 2002, major Y DNA signatures based on markers other than SNPs were recognized. In Western Europe the STR haplotype known as the Atlantic Modal Haplotype was found to be most common by Wilson et al.[6] Even earlier research using RFLP genotyping identified two distinct haplotypes within R-M269. In southeast Europe and southwest Asia (e.g., the Balkans, Georgia and Turkey) "haplotype 35" or "ht35" was found to be a common form of R-M269, whereas in western Europe "haplotype 15" or "ht15" dominated in frequency.[4]

R1b is a sub-clade within the much larger Eurasian MNOPS "macro-haplogroup", which is one of the predominant groupings of all the rest of human male lines outside of Africa, and this whole group, along indeed with all of macro-haplogroup F, is believed to have originated in Asia.

The point of origin of R1b is thought to lie in Eurasia, most likely in Western Asia.[7] T. Karafet et al. estimated the age of R1, the parent of R1b, as 18,500 years before present.[1]

Early research focused upon Europe. In 2000 Ornella Semino and colleagues argued that R1b had been in Europe before the end of the Ice Age, and had spread north from an Iberian refuge after the Last Glacial Maximum.[8] Age estimates of R1b in Europe have steadily decreased in more recent studies, at least concerning the majority of R1b, with more recent studies suggesting a Neolithic age or younger.[7][9][10][11] Only Morelli et al. have recently attempted to defend a Palaeolithic origin for R1b1b2.[12] Irrespective of STR coalescence calculations, Chikhi et al. pointed out that the timing of molecular divergences does not coincide with population splits; the TMRCA of haplogroup R1b (whether in the Palaeolithic or Neolithic) dates to its point of origin somewhere in Eurasia, and not its arrival in western Europe [1]. However, Michael R. Maglio argues that the closest branch of R1b is from Iberia and its small subclades found in West Asia, the Near East and Africa are examples of back migration, and not of its origin.[13]

Barbara Arredi and colleagues were the first to point out that the distribution of R1b STRvariance in Europe forms a cline from east to west, which is more consistent with an entry into Europe from Western Asia with the spread of farming.[11] A 2009 paper by Chiaroni et al. added to this perspective by using R1b as an example of a wave haplogroup distribution, in this case from east to west.[14] The proposal of a southeastern origin of R1b were supported by three detailed studies based on large datasets published in 2010. These detected that the earliest subclades of R1b are found in western Asia and the most recent in western Europe.[7][9][15] While age estimates in these articles are all more recent than the Last Glacial Maximum, all mention the Neolithic, when farming was introduced to Europe from the Middle East as a possible candidate period. Myres et al. (August 2010), and Cruciani et al. (August 2010) both remained undecided on the exact dating of the migration or migrations responsible for this distribution, not ruling out migrations as early as the Mesolithic or as late as Hallstatt but more probably Late Neolithic.[7] They noted that direct evidence from ancient DNA may be needed to resolve these gene flows.[7] Lee et al. (May 2012) analysed the ancient DNA of human remains from the Late Neolithic Bell Beaker site of Kromsdorf, Germany identifying two males as belonging to the Y haplogroup R1b.[16] Analysis of ancient Y DNA from the remains of populations derived from early Neolithic settlements such as the Mediterranean Cardium and Central and North European LBK settlements have found an absence of males belonging to haplogroup R1b.[17][18] It has thus been suggested that the arrival of Y Haplogroup R1b coincided with the arrival of the Centum branch of the Indo-European languages in Western Europe.

R1b* (that is R1b with no subsequent distinguishing SNP mutations) is extremely rare. The only population yet recorded with a definite significant proportion of R1b* are the Kurds of southeastern Kazakhstan with 13%.[7] However, more recently, a large study of Y-chromosome variation in Iran, revealed R1b* as high as 4.3% among Persian sub-populations.[19] In a study of Jordan it was found that no less than 20 out of all 146 men tested (13.7%), including most notably 20 out of 45 men tested from the Dead Sea area, were positive for M173 (R1) but negative for P25 and M269, mentioned above, as well as the R1a markers SRY10831.2 and M17, a study indicates that they are all R1b2-v88 [2].[20] Hassan et al. (2008) found an equally surprising 14 out of 26 (54%) of Sudanese Fulani who were M173+ and P25-.[21] Wood et al. report 2 Egyptian cases of R1-M173 which were negative for SRY10831 (R1a1) and P25 (R1b1), out of a sample of 1,122 males from African countries, including 92 from Egypt.[22] Such cases could possibly be either R1b* (R-M343*) or R1a* (R-M420*) (demonstrating the importance of checking exact mutations tested when comparing findings in this field).

It is also possible that some of the rare examples represent a reversion of marker P25 from a positive back to a negative ancestral state.[23]

Europe Y-DNA. Principle puzzles - highlighted areas where the frequency of haplogroups represent more than a third of the gene pool (> 35%)

An up-to-date compilation of data taking the latest information into account can be found in Cruciani et al. (2010) which can be summarised as follows.[24] As will be discussed below, however, in some parts of western and northwestern Europe, R-M269 frequencies can reach even higher levels.

R1b1*, like R1b* is rare. As mentioned above, examples are described in older articles, for example two in a sample from Turkey,[4] but most cases, especially in Africa, are now thought to be mostly in the more recently discovered sub-clade R-V88 (see below). Most or all examples of R1b therefore fall into subclades R1b1c (R-V88) or R1b1a (R-P297). Cruciani et al. in the large 2010 study found 3 cases amongst 1173 Italians, 1 out of 328 West Asians and 1 out of 156 East Asians.[24] Varzari found 3 cases in the Ukraine, in a study of 322 people from the Dniester-Carpathian region, who were P25 positive, but M269 negative.[25] Cases from older studies are mainly from Africa, the Middle East or Mediterranean, and are discussed below as probable cases of R1b1c (R-V88).

R1b1a is defined by the presence of SNP marker P297. In 2008 this polymorphism was recognised to combine M73 and M269 into one R1b1a cluster.[1] The majority of Eurasian R1b is within this clade, representing a very large modern population. Although P297 itself has not yet been much tested for, the same population has been relatively well studied in terms of other markers. Therefore the branching within this clade can be explained in relatively high detail below.

Poor SNP typing has caused confusion as to whether this group is R1b or Q, for example in Behar et al 5 former cases of R1b-M73 were relabeled as a subclade of PQR2xR1 , suggesting that this group may really be Q or less likely R2, as the so called R1b-m73 samples were negative for the mutation that defined R1.[26] m73 (2011 name) is defined by the presence of SNP marker M73. It has been found at generally low frequencies throughout central Eurasia,[27] and in Altay,[28] but has been found with relatively high frequency among particular populations there including Hazaras in Pakistan (8/25 = 32%);[29] and Bashkirs in Bashkortostan (62/471 = 13.2%), 44 of these being found among the 80 tested Bashkirs of the Abzelilovsky District in the Republic of Bashkortostan (55.0%).[30] Four M73 men were also found in a 523-person study of Turkey,[4] and one person in a 168-person study of Crete.[31]

In 2007, Myres et al. report that out of 193 M73 men found amongst 10,355 widespread men, "all except two Russians occurred outside Europe, either in the Caucasus, Turkey, the Circum-Uralic and North Pakistan regions."[36]

R1b1a2 (2011 name) is defined by the presence of SNP marker M269. R1b1a2* or M269(xL23) is found at highest frequency in the central Balkans notably Kosovo with 7.9%, Macedonia 5.1% and Serbia 4.4%.[7] Kosovo is notable in having a high percentage of descendant L23* or L23(xM412) at 11.4% unlike most other areas with significant percentages of M269* and L23* except for Poland with 2.4% and 9.5% and the Bashkirs of southeast Bashkortostan with 2.4% and 32.2% respectively.[7] Notably this Bashkir population also has a high percentage of M269 sister branch M73 at 23.4%.[7] Five individuals out of 110 tested in the Ararat Valley, Armenia belonged to R1b1a2* and 36 to L23*, with none belonging to known subclades of L23.[37]

European R1b is dominated by R-M269. It has been found at generally low frequencies throughout central Eurasia,[27] but with relatively high frequency among Bashkirs of the Perm Region (84.0%) and Baymaksky Region (81.0%).[30] This marker is present in China and India at frequencies of less than one percent. The table below lists in more detail the frequencies of M269 in regions in Asia, Europe, and Africa.

From 2003 to 2005 what is now R1b1a2 was designated R1b3. From 2005 to 2008 it was R1b1c. From 2008 to 2011 it was R1b1b2.

M269

still un-defined

R-M269* (R1b1a2*)

L23

still un-defined

R-L23* (R1b1a2a*)

L150

still un-defined

R-L150* (R1b1a2a1*)

L51/M412

still un-defined

R-L51*/R-M412* (R1b1a2a1a*)

P310/L11

still un-defined

R-P310/L11* (R1b1a2a1a1*)

U106

R-U106 (R1b1a2a1a1a)

P312

R-P312 (R1b1a2a1a1b)

R-L277 (R1b1a1a1b)

As discussed above, in articles published around 2000 it was proposed that this clade had been in Europe before the last Ice Age,[41] but by 2010 more recent periods such as the European Neolithic have become the focus of proposals. A range of newer estimates for R1b1b2, or at least its dominant parts in Europe, are from 4,000 to a maximum of about 10,000 years ago, and looking in more detail is seen as suggesting a migration from Western Asia via southeastern Europe.[2][7][11][15] Western European R1b is dominated by R-P310.[2]

In this period between 2000 and 2010 that it became clear that especially Western European R1b is dominated by specific sub-clades of R-M269 (with some small amounts of other types found in areas such as Sardinia[7][12]). Within Europe, R-M269 is dominated by R-M412, also known as R-L51, which according to Myres et al. (2010) is "virtually absent in the Near East, the Caucasus and West Asia." This Western European population is further divided between R-P312/S116 and R-U106/S21, which appear to spread from the western and eastern Rhine river basin respectively. Myres et al. note further that concerning its closest relatives, in R-L23*, that it is "instructive" that these are often more than 10% of the population in the Caucasus, Turkey, and some southeast European and circum-Uralic populations. In Western Europe it is present but in generally much lower levels apart from "an instance of 27% in Switzerland's Upper Rhone Valley."[7] In addition, the sub-clade distribution map, Figure 1h titled "L11(xU106,S116)", in Myres et al. shows that R-P310/L11* (or as yet undefined subclades of R-P310/L11) occurs only in frequencies greater than 10% in Central England with surrounding areas of England and Wales having lower frequencies.[7] This R-P310/L11* is almost non-existent in the rest of Eurasia and North Africa with the exception of coastal lands fringing the western and southern Baltic (reaching 10% in Eastern Denmark and 6% in northern Poland) and in Eastern Switzerland and surrounds.[7]

In 2009, DNA extracted from the femur bones of 6 skeletons in an early-medieval burial place in Ergolding (Bavaria, Germany) dated to around 670 AD yielded the following results: 4 were found to be haplogroup R1b with the closest matches in modern populations of Germany, Ireland and the USA while 2 were in Haplogroup G2a.[42]

Population studies which test for M269 have become more common in recent years, while in earlier studies men in this haplogroup are only visible in the data by extrapolation of what is likely. The following gives a summary of most of the studies which specifically tested for M269, showing its distribution (as a percentage of total population) in Europe, North Africa, the Middle East and Central Asia as far as China and Nepal.

This subclade is defined by the presence of the SNP U106, also known as S21 and M405.[2][57] It appears to represent over 25% of R1b in Europe.[2]

U106/S21

still un-defined

R-U106* (R-U106-*)

U198

R-M467/S29/U198 (R-U106-1)

DYS439(null)/L1/S26

R-L1/S26 (R-U106-3)

L48

R-L48* (R-U106-4*)

L47

R-L47* (R-U106-4a*)

L44

R-L44* (R-U106-4a1*)

R-L45 and L46 and L146 (R-U106-4a1a1)

R-Z159 (R-U106-4a2)

L148

R-L148 (R-U106-4b)

L257

R-L257 (R-U106-8)

While this sub-clade of R1b is frequently discussed amongst genetic genealogists, the following table represents the peer-reviewed findings published so far in the 2007 articles of Myres et al. and Sims et al.[36][57]

Along with R-U106, R-P312 is one of the most common types of R1b1a2 (R-M269) in Europe. Also known as S116, it has been the subject of significant study concerning its sub-clades, and some of the ones recognized by the ISOGG website are summarized in the following table.[2] Myres et al. described it distributing from the west of the Rhine basin.[7]

Amongst these, scientific publications have given interpretation and comment on several:-

R-P312-2 (R-Z196) unites several branches of R-P312:-

R-P312-2a (R-M153)is defined by the presence of the marker M153. It has been found mostly in Basques and Gascons, among whom it represents a sizeable fraction of the Y-DNA pool,[54][58] though is also found occasionally among Iberians in general. The first time it was located (Bosch 2001[59]) it was described as H102 and included 7 Basques and one Andalusian.

R-P312-2b (R-L176.2) This subclade is defined by the presence of the marker L176.2. It contains the following:

This subclade is defined by the presence of the marker M167, also known as SRY2627. The first author to test for this marker (long before current haplogroup nomenclature existed) was Hurles in 1999, who tested 1158 men in various populations.[60] He found it relatively common among Basques (13/117: 11%) and Catalans (7/32: 22%). Other occurrences were found among other French, British, Spaniards, Béarnais, and Germans.

In 2000 Rosser et al., in a study which tested 3616 men in various populations[61] also tested for that same marker, naming the haplogroup Hg22, and again it was found mainly among Basques (19%), in lower frequencies among French (5%), Bavarians (3%), Spaniards (2%), Southern Portuguese (2%), and in single occurrences among Romanians, Slovenians, Dutch, Belgians and English.::In 2001 Bosch described this marker as H103, in 5 Basques and 5 Catalans.[59] Further regional studies have located it in significant amounts in Asturias, Cantabria and Galicia, as well as again among Basques.[59] Cases in the Azores have been reported.[citation needed] In 2008 two research papers by López-Parra[58] and Adams,[54] respectively, confirmed a strong association with all or most of the Pyrenees and Eastern Iberia.

In a larger study of Portugal in 2006, with 657 men tested, Beleza et al. confirmed similar low levels in all the major regions, from 1.5%-3.5%.[62]

R-P312-2b2 (R-L165). This subclade is defined by the presence of the marker S68, also known as L165. It is found in England, Scandinavia, and Scotland (in this country it is mostly found in the Northern Isles and Western Isles). It has been suggested, therefore, that it arrived in the British Isles with Vikings.[63]

R-P312-3 (R-U152) is defined by the presence of the marker U152, also called S28.[2] Its discovery was announced in 2005 by EthnoAncestry[64] and subsequently identified independently by Sims et al. (2007).[57] Myres et al. report this clade "is most frequent (20-44%) in Switzerland, Italy, France and Western Poland, with additional instances exceeding 15% in some regions of England and Germany."[36] Similarly Cruciani et al. reported frequency peaks in Northern Italy and France.[65] Out of a sample of 135 men in Tyrol, Austria, 9 tested positive for U152/S28.[66] Far removed from this apparent core area, Myres et al. also mention a sub-population in north Bashkortostan where 71% of 70 men tested were in R-U152. They propose this to be the result of an isolated founder effect.[7]

R-P312-4 (R-L21) is defined by the presence of the marker L21, also referred to as M529 and S145.[2] Myres et al. report it is most common in England and Ireland (25-50% of the whole male population).[7] Known sub-clades include the following:-

R-P312-4b (R-M222). This subclade within R-L21 is defined by the presence of the marker M222. It is particularly associated with male lines which are Irish or Scottish, but especially northern Irish. In this case, the relatively high frequency of this specific subclade among the population of certain counties in northwestern Ireland may be due to positive social selection, as it is suggested to have been the Y-chromosome haplogroup of the Uí Néill dynastic kindred of ancient Ireland.[43] However, it is not restricted to the Uí Néill as it is associated with the closely related Connachta dynasties, the Uí Briúin and Uí Fiachrach.[67] M222 is also found as a substantial proportion of the population of Scotland which may indicate substantial settlement from northern Ireland or at least links to it.[43][68] Those areas settled by large numbers of Irish and Scottish emigrants such as North America have a substantial percentage of M222.[43]

R-P312-4g (R-L193). This subclade within R-L21 is defined by the presence of the marker L193. Many surnames with this marker are associated geographically with the western "Border Region" of Scotland. A few other surnames have a Highland association. R-L193 is a relatively young subclade likely born within the last 2000 years.

R-P312-4h (R-L226). This subclade within R-L21 is defined by the presence of the marker L226, also known as S168. Commonly referred to as Irish Type III, it is concentrated in central western Ireland and associated with the Dál gCais kindred.[70]

R-DF21. This subclade within R-L21 is defined by the presence of the marker DF21 aka S192. It makes up about 10% of all L21 men and is c.3000 years old.[71]

R1b1b is defined by the presence of SNP marker M335. This haplogroup was created by the 2008 reorganisation of nomenclature and should not be confused with R1b1b2, which was previously called R1b1c. Its position in relation to the much more populous sub-clade R1b1a is uncertain.[1] The M335 marker was first published in 2004, when one example was discovered in Turkey, which was classified at that time as R1b4.[4]

R1b1c (formerly R1b1a) is defined by the presence of SNP marker V88, the discovery of which was announced in 2010 by Cruciani et al.[24] The V88 group in Africa may be linked with Chadic languages and an Iberian origin.[72] Apart from individuals in southern Europe and Western Asia, the majority of R-V88 was found in northern and central Africa:

Region

Population

Country

Language

N

Total%

R1b1c (R-V88)

R1b1a2 (R-M269)

R1b1c* (R-V88*)

R1b1c4 (R-V69)

N Africa

Composite

Morocco

AA

338

0.0%

0.3%

0.6%

0.3%

0.0%

N Africa

Mozabite Berbers

Algeria

AA/Berber

67

3.0%

3.0%

0.0%

3.0%

0.0%

N Africa

Northern Egyptians

Egypt

AA/Semitic

49

6.1%

4.1%

2.0%

4.1%

0.0%

N Africa

Berbers from Siwa

Egypt

AA/Berber

93

28.0%

26.9%

1.1%

23.7%

3.2%

N Africa

Baharia

Egypt

AA/Semitic

41

7.3%

4.9%

2.4%

0.0%

4.9%

N Africa

Gurna Oasis

Egypt

AA/Semitic

34

0.0%

0.0%

0.0%

0.0%

0.0%

N Africa

Southern Egyptians

Egypt

AA/Semitic

69

5.8%

5.8%

0.0%

2.9%

2.9%

C Africa

Songhai

Niger

NS/Songhai

10

0.0%

0.0%

0.0%

0.0%

0.0%

C Africa

Fulbe

Niger

NC/Atlantic

7

14.3%

14.3%

0.0%

14.3%

0.0%

C Africa

Tuareg

Niger

AA/Berber

22

4.5%

4.5%

0.0%

4.5%

0.0%

C Africa

Ngambai

Chad

NS/Sudanic

11

9.1%

9.1%

0.0%

9.1%

0.0%

C Africa

Hausa

Nigeria (North)

AA/Chadic

10

20.0%

20.0%

0.0%

20.0%

0.0%

C Africa

Fulbe

Nigeria (North)

NC/Atlantic

32

0.0%

0.0%

0.0%

0.0%

0.0%

C Africa

Yorubad

Nigeria (South)

NC/Defoid

21

4.8%

4.8%

0.0%

4.8%

0.0%

C Africa

Ouldeme

Cameroon (Nth)

AA/Chadic

22

95.5%

95.5%

0.0%

95.5%

0.0%

C Africa

Mada

Cameroon (Nth)

AA/Chadic

17

82.4%

82.4%

0.0%

76.5%

5.9%

C Africa

Mafa

Cameroon (Nth)

AA/Chadic

8

87.5%

87.5%

0.0%

25.0%

62.5%

C Africa

Guiziga

Cameroon (Nth)

AA/Chadic

9

77.8%

77.8%

0.0%

22.2%

55.6%

C Africa

Daba

Cameroon (Nth)

AA/Chadic

19

42.1%

42.1%

0.0%

36.8%

5.3%

C Africa

Guidar

Cameroon (Nth)

AA/Chadic

9

66.7%

66.7%

0.0%

22.2%

44.4%

C Africa

Massa

Cameroon (Nth)

AA/Chadic

7

28.6%

28.6%

0.0%

14.3%

14.3%

C Africa

Other Chadic

Cameroon (Nth)

AA/Chadic

4

75.0%

75.0%

0.0%

25.0%

50.0%

C Africa

Shuwa Arabs

Cameroon (Nth)

AA/Semitic

5

40.0%

40.0%

0.0%

40.0%

0.0%

C Africa

Kanuri

Cameroon (Nth)

NS/Saharan

7

14.3%

14.3%

0.0%

14.3%

0.0%

C Africa

Foulbe

Cameroon (Nth)

NC/Atlantic

18

11.1%

11.1%

0.0%

5.6%

5.6%

C Africa

Moundang

Cameroon (Nth)

NC/Adamawa

21

66.7%

66.7%

0.0%

14.3%

52.4%

C Africa

Fali

Cameroon (Nth)

NC/Adamawa

48

20.8%

20.8%

0.0%

10.4%

10.4%

C Africa

Tali

Cameroon (Nth)

NC/Adamawa

22

9.1%

9.1%

0.0%

4.5%

4.5%

C Africa

Mboum

Cameroon (Nth)

NC/Adamawa

9

0.0%

0.0%

0.0%

0.0%

0.0%

C Africa

Composite

Cameroon (Sth)

NC/Bantu

90

0.0%

1.1%

0.0%

1.1%

0.0%

C Africa

Biaka Pygmies

CAR

NC/Bantu

33

0.0%

0.0%

0.0%

0.0%

0.0%

W Africa

Composite

—

123

0.0%

0.0%

0.0%

0.0%

0.0%

E Africa

Composite

—

442

0.0%

0.0%

0.0%

0.0%

0.0%

S Africa

Composite

—

105

0.0%

0.0%

0.0%

0.0%

0.0%

TOTAL

1822

M343

still un-defined

R-M343* (R1b*)

P25

still un-defined

R-P25* (R1b1*)

V88

still un-defined

R-V88* (R1b1c*)

M18

R-M18 (R1b1c1)

V8

R-V8 (R1b1c2)

V35

R-V35 (R1b1c3)

V69

R-V69 (R1b1c4)

R-P297 (R1b1a)

As can be seen in the above data table, R1b1c is found in northern Cameroon in west central Africa at a very high frequency, where it is considered to be caused by a pre-Islamic movement of people from Eurasia.[22][73]

Suggestive results from other studies which did not test for the full range of new markers discovered by Cruciani et al. have been reported, which might be in R-V88.

Wood et al. reported high frequencies of men who were P25 positive and M269 negative, amongst the same north Cameroon area where Cruciani et al. reported high R-V88 levels. However, they also found such cases amongst 3% (1/32) of Fante from Ghana, 9% (1/11) of Bassa from southern Cameroon, 4% (1/24) of Herero from Namibia, 5% (1/22) of Ambo from Namibia, 4% (4/92) of Egyptians, and 4% (1/28) of Tunisians.[22]

Luis et al. found the following cases of men M173 positive (R1), but negative for M73 (R1b1b1), M269 (R1b1b2), M18 (R1b1a1, a clade with V88, M18 having been discovered before V88) and M17 (R1a1a): 1 of 121 Omanis, 3 of 147 Egyptians, 2 of 14 Bantu from southern Cameroon, and 1 of 69 Hutu from Rwanda.[74]

Pereira et al. (2010) in a study of several Saharan Tuareg populations, found one third of 31 men tested from near Tanut in Niger to be in R1b.[75]

Y-chromosomes known as R1b-M73 first appeared in the literature of Sengupta et al. in 2006 among 5 out of 25 Hazaras.[78] R1b-m73 was the sole type of R1b found in the Kumandin and a dominant clade in the South eastern Bashkirs. However, a 2010 study by Behar et al. showed that these same 5 Hazara did not belong to R1b but to R2 or Q, if this study by Behar is right than the Kumandins have no R1b. Ironically the Bashkirs still have a high percent of R1b (mostly-u152) even though some of the southern Bashkirs had mislabled Q which appeared to be R1b due to confusion between Q1a-m25 and R1b-p25. If Behar et al. is right about m73, then the Southeastern Bashkirs have both R1b and Q while all the other Bashkirs have a high frequency of R1b while only having 1-2% of Q.[79][80]